SCIENCE CHINA

Earth Sciences
• RESEARCH PAPER • Julyy 2010 Vol.53 No.7: 956–963
doi: 10.1007/s11430-010-4001-4

New materials of the steppe mammoth, Mammuthus trogontherii,

with discussion on the origin and evolutionary patterns
of mammoths
WEI GuangBiao1,2*, HU SongMei3, YU KeFu 4, HOU YaMei 5, LI Xin6, JIN ChangZhu5,
WANG Yuan5, ZHAO JianXin7 & WANG WenHua8
1
Chongqing Three Gorges Institute of Paleoanthropology, China Three Gorges Museum, Chongqing 400015, China;
2
Chongqing Three Gorges Culture and Social Development Research Institute, Chongqing Normal University, Chongqing 400047, China;
3
Shaanxi Provincial Institute of Archaeology, Xi’an 710054, China;
4
South China Sea Institute of Oceanology, Chinese Academy of Sciences, Guangzhou 510301, China;
5
Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China;
6
Chongqing Natural History Museum, Chongqing 400013, China;
7
Radiogenic Isotope Laboratory, Centre for Microscopy and Microanalysis (CMM), The University of Queensland, Qld 4072, Australia;
8
Production & Technology Department, Strip Mine, Zalainuoer Coal Industry Co. Ltd., Manzhouli 021412, China

Received May 12, 2009; accepted November 6, 2009; published online June 9, 2010

Recently found materials indicate that the steppe mammoth, Mammuthus trogontherii, survived in northern China into the late
Pleistocene. East Asia is the key area of mammoth evolution after the initial radiation of early forms out of Africa and into
Eurasia at the beginning of the late Pliocene (c. 3.5–3.0 Ma). M. rumanus, M. meridionalis, M. trogontherii, and M. primige-
nius probably formed a continuous and transitional evolutionary lineage within the pan-Eurasian mammoth radiation in East
Asia. Each speciation event of the Eurasian mammoths was followed by a rapid and large-scale dispersal event: out of East
Asia. Allopatric speciation is the main speciation pattern of Mammuthus. The climatic vacillation was severe and frequent in
East Asia from the early part of early Pleistocene (c. 2.6 Ma) onward, which probably brought about successive speciation in
East Asia and the subsequent dispersal of the mammoths.

Mammuthus trogontherii, Mammuthus sungari, MSE, allopatric speciation, out of East Asia, climate

Citation: Wei G B, Hu S M, Yu K F, et al. New materials of the steppe mammoth, Mammuthus trogontherii, with discussion on the origin and evolutionary
patterns of mammoths. Sci China Earth Sci, 2010, 53: 956–963, doi: 10.1007/s11430-010-4001-4

During the last decade, European scholars have made con-
siderable progress in the study of the origin and evolution of
mammoths. Owing to the successive discoveries of a series
of important fossils, the application of a variety of dating
technologies, the gradual unification of terminology and
measuring method, the pattern and process of mammoth
evolution in Eurasia have generally taken shape. The re-
*Corresponding author (email: elephantfossil@yahoo.com.cn)
search of Chinese mammoth fossils lags relatively behind.
The taxonomy and biostratigraphy of Chinese Elephantidae
fossils have been in a long-term state of chaos, due to the
scarcity of remains, weak chronostratigraphy, and poor un-
derstanding of foreign fossils plus slow communication
with foreign colleagues.
In recent years, some achievements have been gained on
the research of Chinese mammoth remains, and the Plio-
cene-Pleistocene biostratigraphic framework based on

© Science China Press and Springer-Verlag Berlin Heidelberg 2010 earth.scichina.com www.springerlink.com
 WEI GuangBiao, et al. Sci China Earth Sci July (2010) Vol.53 No.7 957

mammoth fossils of northern China has been established. while abrasion surface is apparently convex. The cervical
On the basis of the newly discovered mammoth remains and line is distinctly visible. Midline of crown obviously curves
combined with literature and restudy of the specimens pre- toward the lingual side. Plates are weakly convergent to-
viously reported, this paper further enriches the stratigraphic wards the lingual side in occlusal view. There are only 15
distribution of mammoth fossils from Eurasia, especially preserved plates leading to uncertainty of the number of
China, and explores the origin and evolutionary model for original plates, but the reconstruction of the missing distal
large terrestrial mammals represented by the mammoth. part of the crown suggests a total plate number of about 18–
20 (personal communication with van Essen Hans, 2007).
1 Taxonomy of new materials and status of M. Abrasion surface of this molar bears typical morphologi-
sungari Chow et Chang, 1959 cal characters of Mammuthus as follows: elasmodont cheek
teeth, the enamel layers of the mesial and distal sides paral-
1.1 Material from Weihe River terrace of Gaoling lel with each other; wear figure of some plates typically
County, Shaanxi Province showing the tripartite structure of two lateral lamellar
enamel loops with nearly the same size and rectangle-
Elephantidae Gray, 1821
shaped and one central annular enamel loop (wear figure of
Elephantinae Gray, 1821
Palaeoloxodon plates showing the lateral annular and cen-
Mammuthus Brookes, 1828
tral lamellar structures); the three enamel loops fusing into
Mammuthus trogontherii Pohlig, 1885
one single loop with advanced abrasion, with the mesial
Material: The mesial part of a third right upper molar
(Figure 1).
Specimen No.: SMD: 8.
Locality: The northern bank of the Wei River, Shang-
madu, Gaoling County, Xi’an City, Shaanxi Province,
China, with an altitude of 400 m (34°32′N, 109°04′E).
Horizon: Unclear.
Storage: Shaanxi Provincial Institute of Archaeology.
Geological age: Late Pleistocene.
Dating results: 33.858–24.857 ka BP (Table 1).
Dating laboratory: Radiogenic Isotope Laboratory,
Centre for Microscopy and Microanalysis, University of
Queensland, Australia.
Sample selection and analytical methods: We use
TIMS (thermal ionization mass spectrometric) U-series
technique to date dental samples. At first we manually
cleaned teeth by polishing any weathered surfaces, and then
got about 0.06 g fresh specimen for each sample. The
specimens were ultrasonically cleaned in Milli-Q water,
dried, and then each of the specimens was spiked with
0.03–0.05 g 229Th-233U-236U mixed tracer (0.03–0.05 g), and
totally dissolved in 15.8 mol/L HNO3. Further chemical
treatment and TIMS U-series analytical procedures fol-
lowed those described in refs. [1, 2].
Discussion (see Table 2 for measurements): This
Figure 1 Mammuthus trogontherii (Pohlig). (a) SMD: 8, Right M3,
specimen is the mesial part of a moderately worn right M3 Occlusal view; (b) SMD: 8, Right M3, Labial view, Shangmadu, Gaoling
of large dimension. With medium thickness, plates are cov- County, Xi’an City, Shaanxi Province. (c) ZLNE001, Right M3, Occlusal
ered with thick and uneven cement. In lateral view, this view; (d) ZLNE001, Right M3, Lingual view, Zalainuoer, Manzhouli City,
molar is nearly triangular-shaped without preserved roots Inner Mongolia Autonomous Region.

Table 1 Original data of the TIMS U-series dating

232 230 230 234 Uncorr.

Th Th/ Th/ U/ Corr. 230Th Corr. initial
Sample U (ppm) ±2σ (ppb) ±2σ 232
Th 238
U ±2σ 238
U ±2σ 230
Th age ±2σ age (ka) ±2σ 234
U/238U ±2σ
(ka)
SMD-1 0.5583 0.0004 0.4353 0.006 1452.052 0.37312 0.00385 1.3765 0.0022 33.874 0.410 33.858 0.410 1.4144 0.0024
SMD-2 0.7830 0.0006 0.5576 0.003 1201.165 0.28193 0.00185 1.3664 0.0030 24.872 0.192 24.857 0.192 1.3932 0.0032
958 WEI GuangBiao, et al. Sci China Earth Sci July (2010) Vol.53 No.7

Table 2 Comparative measurements for the upper third molars of the three species of Mammuthus

Species Locality of sample P Lmax (mm) Hmax (mm) W (mm) E (mm) LF HI Source
11–14 228.8–317.1 100.2–141.8 85.6–126.4 2.6-4.1 3.7–6.1 93.8–152.7 [3]
Eurasia 11–15 [4]
M. meridionalis
Upper Valdarno, 11–13 240.0–298.0 85.0–122.0 4.5–5.0 [5]
Italy 12–14 220.0–335.0 104.0–147.0 80.0–123.5 2.5-3.8 4.18–6.41 113.0–147.0 [6]
14–21 213.0–358.0 118.0–218.0 57.0–107.5 1.5-3.0 6.0–8.2 145.3–304.9 [3]
Eurasia 17–22 [4]
M. trogontherii
China x17x–x171/2x 264.0–313 160.0–163.0 89–91 ≥2.0 5.8–7.3 [7]
Süssenborn, Germany 18–23 246.0–400.0 129.0–212.0 73.0-120.0 2.0-3.0 4.0–7.0 143.0–208.0 [5]
M. primigenius Eurasia & N. America 20–27 226.0–285.0 135.0–188.5 68.0-113.0 1.3-2.0 6.5–11.1 164.6–211.8 [3]
SMD: 8 ∞15+
222+ 189+ 115.0 3.0 6.1 164.3+
(Xi’an) (about 18–20)
Present specimen
ZLNE001
x19x 280+ 212 104 2.4 6.6 203.8
(Zalainuoer)

and distal sides of the loop parallel with each other; the dis-
tally and/or mesially prominent, small and pointed structure
(loxodont sinus) absent at the central part of plate.
In this specimen, it is the type of enamel fold that most of
all indicates points to the direction of a mammoth. In Pa-
laeoloxodon specimens, the enamel fold is more intense and
at the same time more regular. Especially deep loops (am-
plitude more than twice the enamel thickness) occur both at
the median line and from there at regular intervals in buccal
and lingual direction. In mammoths, there seems to be a
kind of “hesitation” in the enamel to commit to a direction
of folding. In this specimen, for example, in certain places
there is very little enamel folding, and in others it may be
stronger, but it is never as sharp and regular as in Palaeol-
oxodon.
This specimen must be assigned to the genus Mammut-
hus. All the measurements of this molar, such as plate
number (P), lamellar frequency (LF), enamel thickness (E),
and width (W) and height (H) of crown, fall within the
ranges of M. trogontherii (Table 2). The above characters
preclude the possibility of referring the M3 to M. primigen-
ius and M. meridionalis.
U-series dating results show that this specimen is proba-
bly the latest record of M. trogontherii anywhere in the
world. Previously, the latest global record of M. tro-
gontherii was from Marsworth, England with the date 200－
150 ka [4, 8].
1.2 Material from Zalainuoer, Inner Mongolia
Elephantidae Gray, 1821
Elephantinae Gray, 1821
Mammuthus Brookes, 1828
Mammuthus trogontherii Pohlig, 1885
Material: The complete third right upper molar (Figure
1).
Specimen No.: ZLNE001.
Locality: Lingquan Strip Mine, Zalainuoer, Manzhouli
City, Inner Mongolia Autonomous Region (49°20′N,
117°35′E).
Horizon: Bottom of the alluvium of the Zalainuoer old
riverbed.
Storage: Zalainuoer History and Culture Museum.
Geological age: Late Pleistocene.
Absolute age: c. 33.7 ka BP (AMS 14C) [9].
Discussion (measurements see Table 2): Similar to the
above specimen, abrasion surface of this molar bears typical
morphological characters of Mammuthus. The enamel lay-
ers of the mesial and distal sides are parallel with each other
in medium or advanced wear. The labio-lingual length of
the central enamel loop is nearly equal to those of the two
lateral enamel loops (in Palaeoloxodon, the central part is
rather wider mesio-distally than the lateral parts, showing
an elongated rhombic shape). A prominent and obtuse me-
sial and/or distal median sinus is developed in the central
part (in Palaeoloxodon, a small and acute loxodont sinus is
visible in the same position). There are irregular folds of
enamel layer in this specimen (viz., strong folds in places,
little folding elsewhere). This specimen should be referred
to the genus Mammuthus for all the measurements of the
molar, especially plate number, fall within the range of this
species.
As the type materials of M. sungari Chow et Chang,
1959 (mammoth skeletons unearthed from Zalainuoer, Inner
Mongolia Autonomous Region), are well-known, M. sun-
gari is till employed by Chinese scholars for local Mam-
muthus. However, we consider that the specific name is a
synonym of M. trogontherii for the following reasons.
Firstly, when Zhou and Zhang [10] erected M. sungari,
there were only referred specimens without a holotype,
which violated paleontological nomenclatural rules. Mean-
while, the referred specimens, most of which were M1/m1,
M2/m2 and P/p, were very few and lacked skull material or
complete M3/m3, which are most significant for identifying
mammoths at the specific level. The only M3 (V. 2053) was
broken, without distinct characters.
Secondly, the formal diagnosis of M. sungari is very am-
biguous and it could not be distinguished qualitatively or
quantitatively from other species of Mammuthus.
Further, the typical materials of M. sungari assigned by
 WEI GuangBiao, et al. Sci China Earth Sci
Chow and other researchers, after restudy, can be shown to
be referred to M. trogontherii (such as one third left m3
from Ji’xian County, Heilongjiang Province [11]) or M.
primigenius (such as the skeleton from Zhaoyuan County,
Heilongjiang Province [12]) respectively according to mor-
phological characters and measurements.
Lastly, the general acceptance of the specific name M.
sungari by most of the Chinese scholars mainly depends on
its gigantic stature and the more primitive morphological
characters compared with M. primigenius specimens; these
two points are just the main characters of M. trogontherii.
To date, three mounted skeletons have been considered
to represent M. sungari, i.e., the skeletons of No.1, No.2 and
No.3 from Zalainuoer. The first two are exhibited in Inner
Mongolia Museum whereas the last one is conserved in the
Zalainuoer History and Culture Museum. There is also a
mounted mammoth skeleton from Zhaoyuan, Heilongjiang,
preserved in Heilongjiang Provincial Museum. Compared
with the mixed mounting of No.1 and No.2 skeletons, the
80% recovery of No.3 skeleton is more credible. The height
of the mounted No.3 skeleton is 4.33 m, which is almost the
same as the height of the large M. trogontherii skeleton
from Europe.
According to Wang Zhengyi of Zalainuoer History and
Culture Museum (personal communication, 2008), the
above M3 is from the same locality and horizon as No.3
mammoth skeleton. Being similar, they are likely the same
species. The measurement of one complete third m3 (with
complete plate number of 18 and enamel thickness of 3 mm)
[13] also falls within the range of M. trogontherii. Accord-
ing to the present data, the three skeletons of mammoth, as
well as the above M3, should be referred to M. trogontherii.
The mammoth skeleton from Zhaoyuan, with the height of
3.33 m after mounting (c. 21.2 ka BP) [12], is similar in size
to the remains from NE Siberia and Europe and should be
referred to M. primigenius.
Remains previously assigned to M. sungari can be di-
vided into two morphologies after restudy. The primitive
type represents an advanced remnant group of M. tro-
gontherii, probably the latest record of that species. A de-
rived form represents M. primigenius migrated from NE
Siberia. Relative to the Late Pleistocene M. primigenius
from NE Siberia and Europe, the M. trogontherii from
northern China had more primitive characters and gigantic
stature, and the evolutionary level of the latter is nearly the
same as M. columbi from North America. It is difficult to
judge whether this group should be divided into spatial or
temporal subspecies, or whether M. trogontherii and M.
primigenius overlapped spatially or temporally in northern
China, due to the scarcity of materials and the inadequate
degree of study.
With climate fluctuations and environmental changes, the
remnant group of Chinese M. trogontherii ranged in 35°–
50°N during the late Pleistocene. There was a large-scale
cooling event during 40–30 ka, pushing southward M.
July (2010) Vol.53 No.7 959
primigenius into the Jin’an area [14]. The time interval
largely agrees with the latest record of Chinese M. tro-
gontherii. Consequently, it is most probable that during this
cooling event the latest M. trogontherii gave way to M.
primigenius, which migrated from NE Siberia with the lati-
tudinal shift in steppe environment.
2 Evolution and dispersal patterns of Eurasian
mammoths
2.1 Origin and evolution of species of Mammuthus
The generalized mammoth genus Mammuthus originated
during 5–4 Ma from Africa, where the early Pliocene spe-
cies M. subplanifrons Osborn, 1928 and the late Plio-
cene-early Pleistocene species M. africanavus Arambourg,
1952 were named [3]. The former taxon incorporates the
earliest known mammoth materials living in African tropi-
cal circumstances and may be the ancestor of all mammoths
flourished in Eurasia and North America. M. africanavus is
considered as the descendant of M. subplanifrons and an
evolutionary “dead end” restricted to Africa [15].
So far, five continental mammoth species are generally
accepted by different authors for Eurasia and North Amer-
ica: M. rumanus Stefanescu, 1924, M. meridionalis Nesti,
1825, M. trogontherii Pohlig, 1885, M. primigenius Blu-
menbach, 1799, and M. columbi Falconer, 1857. In addition,
there are still some small-sized or pygmy mammoth types in
the islands of Eurasia and North America, such as M. pro-
tomammonteus from Japan, M. lamarmorae from Sardinia,
and M. exilis from Channel Islands, California. M. pro-
tomammonteus from Japan is a small-sized mammoth
without dwarfism. This paper emphasizes the four kinds of
Eurasian continental mammoth species.
The earliest Eurasian mammoth fossils are named M.
rumanus because the holotype was found in the Dacic Basin,
Romania. The first emergence of this species is almost at
the same time in both Europe and China, i.e., mammalian
biozone MN16a, correlated to palaeomagnetic data in the
middle Gauss subchron, c. 3.5–3.0 Ma [4, 16]. The first
appearance of M. rumanus in Eurasia follows the first im-
portant dispersal event of mammoth lineage: Out of Africa,
as well as the shift of center of the mammoth evolutionary
lineage. M. rumanus is known to have survived until the
early part of early Pleistocene (c. 2.5 Ma) in Europe [4, 17],
but somewhat earlier in China [7, 16]. Due to the scarcity of
materials, it still needs further study concerning the locus
and time of origin of M. rumanus in Eurasia.
The holotype of M. meridionalis was found in Upper
Valdarno, Italy, situated in southern Europe. The distribu-
tion of M. meridionalis was limited in Eurasia and nearby
islands. The temperate forest environment of Europe during
most of the interval of 2.0–0.7 Ma [15] was advantageous to
the survival and evolution of M. meridionalis. The earliest
European M. meridionalis (M. gromovi, coined by Alexeeva
960 WEI GuangBiao, et al. Sci China Earth Sci
and Garutt, 1965, for remains from the Khapry Faunal
Complex in the south of European Russia) is now dated to
early Pleistocene, early MN17, 2.6–2.2.2 Ma [18] or
2.4–2.2 Ma [8]. The emergence of M. meridionalis in West
Europe was somewhat later than that in East Europe, such
as at the Chilhac site in France with the age of 2.0 Ma [4, 8]
and the Upper Valdarno, Italy with 2.0–1.77 Ma [19].
The most primitive M. meridionalis remains from China,
originally identified as Archidiskodon planifrons by Wang
[20], were found in Houhecun, Dali, Shaanxi, including
relatively complete skull and postcranial material. Accord-
ing to the original plate and description of Wang [20], these
materials are most similar to the typical specimens of M.
meridionalis both from Europe and China. Specifically, the
Houhecun mammoth is probably in a more primitive evolu-
tionary stage than the Khapry mammoth. As the most im-
portant indicator of the evolutionary level of mammoth, the
lowest complete plate number of M3 observed in Khapry
fossils is 12 [4, 17] while that of the Houhecun remains is
only 11 [20]. The age of the Houhecun fauna correlates to
the Matuyama-Gauss palaeomagnetic transition [21], c. 2.6
Ma, which is a little earlier or equal to the age of Khapry
remains.
There are remains of M. rumanus, the precursor of M.
meridionalis, in the Youhe Formation subjacent to the
Houhecun Formation. These two species, without obvious
overlap, display continuity in temporal distribution and
morphological characters [4, 7, 16]. On the contrary, there
has not been any record of M. rumanus remains in the
Khapry district. M. meridionalis most likely originated from
northern China at c. 2.6 Ma and subsequently rapidly spread
westward to Europe with Equus from North America. The
major growth of global ice at about 2.6 Ma occurred, while
the arctic icecap appeared and the Bering Landbridge
formed [21].
Micro-mammal fossils and ostrich eggs from Houhecun
also indicated a dry and cool grassland environment,
whereas the Youhe fauna complex including M. rumanus
suggested a moist and temperate forest environment [20]. It
is probable that just the cooling event provided selection for
the faunal change, also for the origin of M. meridionalis
from an indigenous M. rumanus population. There were
likely environmental differences for M. meridionalis be-
tween east and west. On the one hand, European M. merid-
ionalis adapted to a temperate forest environment and was
replaced by the eastern M. trogontherii at the early part of
middle Pleistocene c. 0.7 Ma [4]. On the other hand, East
Asian M. meridionalis had adapted to dry and cool grass-
land environment around 2.6 Ma. With climatic fluctuations
in East Asia, mammoths gradually adapted to withstand
cold and dry conditions and finally evolved to M. primi-
genius, which possessed long fur and survived in the tundra.
The first emergence of M. meridionalis and Equus in
Eurasia is called the Elephant-Equus event. However,
long-term ambiguity on the definition of “Elephant” caused
July (2010) Vol.53 No.7
controversy on defining the first appearance of “Elephants”
in Eurasia as the beginning of the Quaternary. Generally,
“Elephant” includes members of Elephantinae, such as the
extinct genera of Primelephas, Palaeoloxodon, and Mam-
muthus, as well as the extant genera of Elephas and Loxo-
donta. The mammoth evolutionary lineage in Eurasia is one
of continuous evolution, extensive distribution, and distinct
characters. The first emergence of one new species in the
lineage can be the basis of geological age division. However,
there is temporal and regional asynchronization on biologi-
cal and climatic events, while palaeomagnetic reversal
events are globally simultaneous. Thus, it is preferable to
avoid biological events as the basis of geological age divi-
sion, and we adopt the international dividing standard based
on palaeomagnetic reversal events passed by the Interna-
tional Commission on Stratigraphy (ICS) in May, 2009.
According to this new dividing standard, the Pleistocene/
Pliocene boundary locates at the Matuyama/Gauss reversal
(c. 2.6 Ma).
The earliest global M. trogontherii remains were recov-
ered in situ from Majuangou-3, Nihewan basin, China, with
palaeomagnetic dating of 1.66 Ma, and North China is con-
sidered as the locus of origin of M. trogontherii [22, 23].
Among the steppe mammoth fossils recovered in China, one
complete M3 from Pinglu, Shanxi [11] has only 17 com-
plete plates while other measurements (such as Lamellar
Frequency and Hypsodonty Index) also show more primi-
tiveness than among the Majuangou materials [16]. The M3
from Pinglu most likely represents the earliest and most
primitive steppe mammoth. Unfortunately, there is no exact
locality and horizon record of this molar.
As discussed above, M. trogontherii in China survived to
the late Pleistocene. The steppe mammoth experienced a
large-scale dispersal from northern China during 1.5–1.2
Ma. Firstly, it spread northward to NE Siberia, and then
entered North America via Bering Landbridge where it
evolved to the large-sized local taxon M. columbi, which
became extinct in the late Ice Age, c. 11 ka BP. Secondly, it
migrated eastward to Japan and evolved as an insular spe-
cies M. protomammonteus, which became extinct around
0.5 Ma [4, 24]. Lastly, it dispersed westward to Europe via
the Middle East, and displaced the local M. meridionalis
around 0.7 Ma [4, 8]. In Europe, the replacement of M. me-
ridionalis by M. trogontherii was intertwined with a complex
pattern of stasis, transformation, and hybridization [4, 8].
M. primigenius (wooly mammoth) adapted to dry and
cold environment and originated from M. trogontherii in
NE Siberia as early as 0.8 Ma [8]. The primitive M. primi-
genius spread westward to Europe, eastward to North
America via the Bering Landbridge and southward to China
and Japan. The insular group of M. primigenius survived as
late as 3.7 ka [25]. Based on the present data, the first ap-
pearance of M. primigenius in North China and Northeast
China was probably no earlier than 40 ka. The Chinese M.
primigenius, with continental M. primigenius from Eurasia
 WEI GuangBiao, et al.
and North America together, became extinct at the end of
the Ice Age, c. 11 ka BP.
2.2 Patterns of mammoth evolution
During the late Pliocene and Pleistocene, the mammoth
lineage experienced eight Mammoth Speciation Events
(MSE), resulting in the emergence of five continental spe-
cies (M. rumanus, M. meridionalis, M. trogontherii, M.
primigenius and M. columbi) and three insular species (M.
protomammonteus, M. lamarmorae and M. exilis). The pat-
tern and process are shown in Figure 2. It is probable that
the transformation within M. rumanus, M. meridionalis and
M. trogontherii occurred in northern China whereas the
shift from M. trogontherii to M. primigenius took place in
NE Siberia.
Figure 2 Stratigraphic distribution of Eurasian and North American Mammuthus (excluding insular species).
Sci China Earth Sci July (2010) Vol.53 No.7 961
As the dominant speciation mode for mammoths, allo-
patric speciation can be further divided into peripatric
speciation and vicariance speciation [26]. The appearance of
M. rumanus (MSE1), M. columbi (MSE5), M. protomam-
monteus (MSE6), M. lamarmorae (MSE7) and M. exilis
(MSE8) apparently belongs to the typical vicariance speci-
ation (geographic speciation) whereas the emergence of
M.meridionalis (MSE2), M. trogontherii (MSE3), and M.
primigenius (MSE4) accords with peripatric speciation (see
Figure 2). Recent work has tended to emphasize the power
of local habitat variation, rather than mere isolation, in
driving peripheral populations to speciation via adaptive
natural or sexual selection [4, 27].
As discussed above, the geological distribution of every
species of the genus Mammuthus indicates regional asyn-
chronization (see Figure 3). During the process of mam-
962 WEI GuangBiao, et al.
Figure 3 Model of Eurasian and North American mammoth evolution and dispersal at specific level.
moth evolution, there is large-scale dispersal after each
speciation event. The dispersal route in the late Pliocene
was from Africa to Eurasia (Out of Africa). Later in the
Pleistocene, the mammoth evolution and dispersal center
turned to East Asia including northern China and NE Sibe-
ria. The main dispersal routes of mammoths are from East
Asia westward to Europe and eastward to North America
(Out of East Asia).
The relationship between mammoths and humans de-
serves discussion. The coexistence of mammoth fossils and
human remains or artifacts in many Paleolithic sites of
Eurasia since the Pleistocene indicates that mammoths and
humans inhabited the same biozone for a long time. For
example, the coexistent interval of steppe mammoths and
humans was at least 0.6 Ma in the Nihewan Basin of North
China. The interval between the age of Majuangou site and
Donggutuo site both including M. trogontherii fossils and
artifacts was 1.7–1.1 Ma [16]. At the Majuangou site, one
recovered rib of M. trogontherii with distinct chopping and
scraping traces demonstrates mammoth-hunting by humans.
There is synchronization between mammoth speciation and
human evolution. The emergence of Australopithecus
afarensis, Homo rudolfensis/Homo habilis, Homo erectus,
and Homo heidelbergensis may correlate with the evolution
of M. rumanus (3.5–3.0 Ma), M. meridionalis (2.6–2.5 Ma),
M. trogontherii (1.8 Ma), and M. primigenius (0.8 Ma). On
the other hand, the appearance of new species of mammoth,
as many other terrestrial mega-mammals, closely follows
palaeomagnetic reversal or global climatic events. Detailed
study is required concerning the correlations within the
Sci China Earth Sci July (2010) Vol.53 No.7
above factors.
During the evolution and dispersal of the mammoth
lineage, morphological variations of the skull, mandible,
and molars reflected environmental selection and corre-
sponding adaptation. For example, the evolutionary trends
of molars could be recognized as increase of plate number
and height of crown and decrease of cement thickness,
strengthening the chewing ability. Mammoths from moist
and warm tropical or subtropical forest environments of
Africa adapted to dry and cool grassland environments of
the mid-latitude or cold tundra at high latitude. Corre-
spondingly, the food source was also broadened from arbor
(fruit, bark and leaf) and shrub to herb; the former were rich
in nutrition, easily edible and less wearing on molars while
the latter were poor in nutrition, difficult to chew and more
abrasive.
An increasing number of scholars recognizes that the
evolutionary trend or rate of terrestrial mega-mammals
mainly depends on the changes of climate and ecology. It is
probable that mammoths and humans were able to spread out
of Africa successively or simultaneously just because of the
remarkable climatic and environmental changes after the late
Pliocene. From the data of deep-sea records in the South
China Sea [28] and loess deposit sequence [29], East Asia
experienced frequent and dramatic climate fluctuations since
the early part of early Pleistocene, c. 2.6 Ma. The uplift of
Tibet is probably one important reason of the global climate
deterioration [30]. Climatic fluctuations since the early part of
early Pleistocene probably stimulated successive mammoth
speciation and dispersal events out of East Asia.
 WEI GuangBiao, et al. Sci China Earth Sci
We would like to thank Mr. Wang Zhengyi from Zalainuoer History and
Culture Museum and Mr. Wei Zhengyi from Heilongjiang Provincial Mu- 13
seum for providing us precious fossil specimens and literatures. Many
thanks go to Mr. van Essen Hans from Leiden University, Netherlands for
his kindness in offering the instructive suggestion on specimens identifica- 14
tion. Our thanks are also due to Prof. Qiu Zhanxiang and Prof. Qiguoqin
from Institute of Vertebrate Paleontology and Paleoanthropology, Chinese
Academy of Sciences (IVPP) for improving the draft version of this manu- 15
script and for providing valuable suggestions. We thank Lawrence J. Flynn 16
of Harvard University for his comment and English correction. We are
also grateful of Mrs. Wang Chuanping and Li Xiaolong from China Three
Gorges Museum, Chongqing for their strong support. This study was sup-
ported by Key Knowledge Innovation Project of Chinese Academy of Sci- 17
ences (Grant No. KJCX2-YW-106), and National Basic Research Program
of China (Grant No. 2006CB806400).
18
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